Digitized by the Internet Archive in 2011 with funding from University of Illinois Urbana-Champaign http://www.archive.org/details/morphologytaxono12haye ' / ILLINOIS BIOLOGICAL MONOGRAPHS Volume XII PUBLISHED BY THE UNIVERSITY OF ILLINOIS *, URBANA, ILLINOIS I EDITORIAL COMMITTEE John Theodore Buchholz Fred Wilbur Tanner Charles Zeleny, Chairman S70.S~ XLL '• / IL cop TABLE OF CONTENTS Nos. Pages 1. Morphological Studies of the Genus Cercospora. By Wilhelm Gerhard Solheim 1 2. Morphology, Taxonomy, and Biology of Larval Scarabaeoidea. By William Patrick Hayes 85 3. Sawflies of the Sub-family Dolerinae of America North of Mexico. By Herbert H. Ross 205 4. A Study of Fresh-water Plankton Communities. By Samuel Eddy 321 LIBRARY OF THE UNIVERSITY OF ILLINOIS ILLINOIS BIOLOGICAL MONOGRAPHS Vol. XII April, 1929 No. 2 Editorial Committee Stephen Alfred Forbes Fred Wilbur Tanner Henry Baldwin Ward Published by the University of Illinois under the auspices of the graduate school Distributed June 18. 1930 MORPHOLOGY, TAXONOMY, AND BIOLOGY OF LARVAL SCARABAEOIDEA WITH FIFTEEN PLATES BY WILLIAM PATRICK HAYES Associate Professor of Entomology in the University of Illinois Contribution No. 137 from the Entomological Laboratories of the University of Illinois . T U .V- TABLE OF CONTENTS 7 Introduction Q Economic importance Historical review 11 Taxonomic literature 12 Biological and ecological literature Materials and methods 1%i Acknowledgments Morphology ]* 1 ' The head and its appendages Antennae. 18 Clypeus and labrum ™ 22 EpipharynxEpipharyru Mandibles. Maxillae 37 Hypopharynx <w Labium 40 Thorax and abdomen 40 Segmentation « 41 Setation Radula 41 42 Legs £ Spiracles 43 Anal orifice 44 Organs of stridulation 47 Postembryonic development and biology of the Scarabaeidae Eggs f*' Oviposition preferences 48 Description and length of egg stage 48 Egg burster and hatching Larval development Molting 50 Postembryonic changes ^4 54 Food habits 58 Relative abundance. Hibernation Pupation 63 Length of life cycle Melolonthinae 64 Rutelinae 66 Dynastinae °' 70 Cetoniinae Laparosticti Taxonomy Keys 73 Summary Bibliography Explanation of plates Index 115 91] LARVAL SCARABAEOIDEA—HAYES INTRODUCTION The larvae of the superfamily Scarabaeoidea are commonly called white grubs or grubworms. Forbes, writing in 1891 on insects of this group in relation to their life histories, said that "it is necessary that the observer should learn to distinguish species of these insects in the grub and larval stage." Among the numerous genera, the species of which are usually more readily recognized in the adult form, there are many species that are almost indistinguishable from one another in the larval stage. To identify them it is usually necessary to rear the specimens. This requires a long period of time and often involves dangers of loss from one cause or another during the period of rearing. In a number of groups, especially in the sub- family Melolonthinae, the life-cycle covers a period of from two to three years, and in northern latitudes even four years. Such a long larval stage makes rearing difficult. Moreover, since the larval life is spent under the surface of the soil, it is difficult to maintain rearing conditions as near nor- mal as necessary. Not only do the representatives of the various sub- families differ in length of life in various localities, but they may exhibit wide degrees of variance even in the same localities. Rapid strides have been made in our knowledge of the life histories of these insects within the last twelve years (1917-29), and it is becoming increasingly more important, as we survey the great differences exhibited, that we be able to recognize the larval forms. As with all other groups of insects, our knowledge of the developmental stages lags behind our knowledge of the adults. This is especially true in the fields of morphology and taxonomy and to a lesser extent in the field of biology. Prior to the studies by Davis and others, which have materially increased our knowl- edge of the group, most of our knowledge of the biology of these insects has been based on fragmentary accounts scattered in the literature. The present work deals with morphological, taxonomic, and biological studies of the various species of white grubs. The morphological studies have aided materially in constructing the diagnostic keys to the various genera which are included. Life-history studies made by the writer, some of which have, in part, appeared in different publications, are here brought together and briefly summarized to make the present work as nearly a complete unit as possible. 8 ILLINOIS BIOLOGICAL MONOGRAPHS [92 Economic Importance Of the four families, Passalidae, Lucanidae, Scarabaeidae, and Trogidae, comprising the superfamily Scarabaeoidea (or Lamellicornia), the family Scarabaeidae is the best known and most important. The others have only slight economic importance. The family Passalidae, which inhabits decaying wood, is represented in tropical forests by numerous species but in North America by only one species, Passalus cornutus Fab. Some species of this family are of biological interest because of a certain degree of social organization ascribed to them, the adults being known to attend the larvae and care for them throughout their period of development to maturity. The adults chew the woody food and prepare it before feeding it to the larvae. The hastening of decay in rotten logs can be said to be a beneficial phase exhibited by this group of insects. The Lucanidae, or stag-beetles, exhibit wide sexual differences in the adult stage. Their larvae, like the Passalidae, live in rotten logs or roots. About 300 species of Lucanidae occur throughout the world, but only 30 species are recorded by Leng (1920) from North America. Aside from their biological interest, the unique stridulation structures of the larvae and the benefits incurred from the hastening of decay, the family can be con- sidered of little importance. The family Trogidae is one of our newer families, having recently been raised to its present position from subfamily rank in the Scarabaeidae. Of the genera occurring in North America, the genus Trox is the most important. The larvae of Trox are scavengers and are usually found in dried, decomposing, animal matter or on the soil under such matter. Little is known of their life history. The Scarabaeidae in the adult stages are known as chafers, May beetles, June bugs, dung beetles, and dor beetles. They constitute a large family with about 14,000 species known from all parts of the world. The family is divided into two categories, or groups, the Laparosticti and the Pleurosticti. The Laparosticti are, for the most part, coprophagous in habits. The Coprinae, Aphodinae, and Geotrupinae are the better known subfamilies with coprophagous habits. A few of the Coprinae are known to be myrmecophilous and live as symphiles in the nests of ants. The copro- phagus habit is, by far, the most important. The group Pleurosticti contains a large number of important economic insects. These occur in four sub- families: the Melolonthinae, the Rutelinae, the Dynastinae, and the Cetoniinae. The larvae of the subfamilies here considered are recognized in many parts of the world as pests of planted crops and are almost universally known as "white grubs." One of the most notorious members of the group is the European "Hanneton," Melolontha melolontha, which has been 93] LARVAL SCARABAEOIDEA—HAYES 9 known for over a century as a destructive enemy of the roots of crops. It belongs to the subfamily Melolonthinae, to which our common American species of May beetles, or June bugs, are assigned. Certain species of Phyllophaga (Lachnosterna), both adults and larvae, are perhaps the most destructive members of the family in North America, and their injury is too well known to require extended discussion. In some of the states west of the Mississippi the greatest loss from white grubs occurs where the larvae of P. lanceolata (Say) attack the roots of wheat and often destroy thousands of acres in a single season. Considering the country as a whole, most of the principal crops as well as native prairie pastures, meadows, and lawns have suffered injury from various species. In some localities gardens suffer. Potatoes especially have paid a heavy toll in the last few years. Strawberry beds have been the object of regular attacks, and ornamentals of all kinds, particularly those recently set out, have been seriously damaged. Many lawns have been killed by the grubs, and serious loss has occurred in the bluegrass pastures. According to Davis (1918), alfalfa is usually considered free from injury, but it has been found damaged in a number of localities. In attacking this crop, the grub enters the tap-root an inch or two below the surface of the soil, excavates a cavity, and burrows upward and downward in much the same manner as some of the root-boring insects. Among the Rutelinae, two species, Pelidnota punctata L. and Cotalpa lanigera (L.) are notably injurious in the United States. The damage caused by the spotted grapeleaf beetle, Pelidnota punctata, to the foliage of grapes during some years attracts considerable attention, but the larvae, living in rotten wood, such as logs and old stumps, can be regarded as beneficial because their activities in these situations hasten decay. The goldsmith beetle, Cotalpa lanigera, has been reported by Davis (1916) as ranking next in importance to the species of Phyllophaga. He says that the grubs in Michigan are destructive to raspberry bushes, strawberries, corn, and grass. Other native species of Rutelinae treated here are not of so much economic importance, but Anomala innuba at times becomes numerous. The beetles of this species do some damage to the kernels of growing wheat, which they attack while "in the milk," and the grubs feed on the roots of various plants. Two important imported members of this subfamily are the Japanese beetle, Popilliajaponica Newm., and the Asiatic beetle, Anomala orientalis (Waterhouse).